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A girl named Penny and a dog named Bolt, who has various superpowers, must constantly thwart the evil plans of the nefarious Doctor Calico. However, little does Bolt know he is actually just a star on a hit television series called Bolt. To gain a more realistic performance, the TV show's producers have deceived Bolt his entire life, arranging the filming in such a way that Bolt believes the television show is real and he really has superpowers. After filming completes for the latest episode, Bolt escapes from his on-set trailer when he mistakenly believes Penny has been kidnapped by the television villain. He attempts to break through a window, knocking himself unconscious and falls into a box of styrofoam peanuts. With no one aware Bolt is in the box, it is shipped from Hollywood to New York City. In New York, he meets Mittens, a female alley cat who bullies pigeons out of their food. Bolt, convinced Mittens is an evil nemesis who works for Doctor Calico, forces Mittens to help him get back to Hollywood, and the two start their journey westward on a truck. Meanwhile, in Hollywood, Penny is deeply saddened over Bolt's disappearance but is forced by the studio to continue filming with a Bolt lookalike. As their adventure proceeds, Bolt starts to notice that his superpowers aren't working, and rationalizes this is the effect that styrofoam has on his body.

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